IAWA Journal, Vol. 20 (4), 1999: 441-455

WOOD ANATOMY OF THE TRIBE DIPTERYGEAE WITH COMMENTS ON RELATED PAPILIONOID AND CAESALPINIOID LEGUMINOSAE by Peter Gasson

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom

SUMMARY

The wood anatomy of all three genera in the tribe Dipterygeae, , Pterodon and is described here in a systematic context. Dipteryx and Pterodon have short, narrow, storied rays and fine intervessel pitting, whereas Taralea has narrow non-storied rays and much finer intervessel pitting. The wood of T. casiqui­ arensis is much more similar to that of Dipteryx and Pterodon than to its congeners, and a new combination based on both wood and morphological features is being published in Kew Bulletin (Lewis & Gasson, in press). The Dipterygeae may form an outlying group to the , although its nearest affinity has been thought to be with Monopteryx (Sophoreae). Here, the wood anatomy of the three genera is compared with that of Sophoreae, Swartzieae and Dalbergieae. Key words: Dipterygeae, Dipteryx, Pterodon, Taralea, wood de­ scriptions.

INTRODUCTION

The tribe Dipterygeae (Leguminosae, Papilionoideae) comprises three genera from Central and South America: Dipteryx, Taralea and Pterodon. This tribe may form an outlying group to the Dalbergieae, although Polhill (l981b) considered the nearest affinity to be with Monopteryx in the Dussia group in the tribe Sophoreae (Polhill 1981a). In this paper, the wood anatomy of the three genera is described and com­ pared with that of Sophoreae (Gasson 1994, Fujii et al. 1994), Swartzieae (Cowan 1981; Gasson 1996; Gasson & Webley 1999), and most genera of Dalbergieae. This work is part of a systematic study of the wood of all available genera in the Papil­ ionoideae, and will be followed by a survey of Dalbergieae wood anatomy. All three genera of Dipterygeae have previously been described separately in wood identification manuals, but there are some discrepancies between anatomy and that are highlighted and discussed in this paper.

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MATERIALS AND METHODS

The descriptions are based on sections of the following wood sampIes: Dipteryx Schreb.: D. alata: SJRw 47807, Bolivia; SJRw 50210, Bolivia - D. ferrea: MADw 12465, Brazil; (D. cf. ferrea) MADw 22287, Peru- - D. lacunifera: MADw 45211, Brazil - D. magnifica: MADw 46441, Brazil; SJRw 22606 - D. micrantha: SJRw 36652, Brazil - D. panamensis: SJRw 45644, Panama; SJRw 3015 - D. polyphylla: SJRw 33835, Brazil; SJRw 42406, Venezuela - D. punctata: SJRw 34090, Brazil; SJRw 43591, Guyana; MADw 31887, Venezuela - D. rosea (probably a syno­ nym of D. odorata): MADw 31888, Venezuela. Pterodon Vogel: P abruptus: B.B. Klitgaard 75, Brazil - P emarginatus (labelIed P pubescens): MADw 18849, Bo1ivia. Taralea AubI.: T casiquiarensis: SJRw 42161, Venezuela - T crassifolia (labelIed D. crassifolia): MADw 41847 (= Uw 27320), Guyana - T oppositifolia: SJRw 23633, Brazil; SJRw 36631, Brazil; SJRw 36655, Brazil- T rigida (labelIed D, rigida): MADw 45218, Brazil; SJRw 54508, Venezuela.

The names given here have been checked by Gwilym Lewis to ensure that the speci­ mens have been referred to the correct and .

GENERAL WOOD DESCRIPTION

Growth rings not seen. Wood diffuse-porous. Vessels solitary, in pairs, some short ra­ dial multiples and clusters, but in no predominant pattern. Vessel elements relatively short, with alternate, fine (but very fine in Taralea species except for T. casiquiarensis), vestured intervascular pits and simple perforation plates. Vessel-ray pitting similar to intervascular pitting in size and shape. Fibres thick- to very thick-walled, non-septate with slit-like, apparently simple pits. Axial parenchyma scanty paratracheal to vasi­ centric, aliform and confluent, storied except in most Taralea species. Rays uniseriate or biseriate, three cells wide in a few species, mainly homocellular, consisting mostly of procumbent cells, storied in Dipteryx, Pterodon and T. casiquiarensis, but not storied in the other Taralea species. Prismatic crystals in chambered axial parenchyma cells present in most species, possibly also in some fibres adjacent to axial parenchyma.

GENERIC WOOD DESCRIPTIONS AND PREVIOUS LITERATURE

Dipteryx - Fig. 1-17 About 10 species from Central and South America, nine examined here. There are many publications with wood anatomical descriptions, including Anonymous (1981 a, 1981b), Pfeiffer (1926), Record & Mell (1924), Lindeman et al. (1963), Sudo (1963), Reinders Gouwentak & Rijsdijk (1968), Huizzi (1974), Mainieri (1978), Dechamps (1980), Gaiotti de Peralta & Edlmann Abbate (1981), Detienne et al. (1982), Detienne & Jacquet (1983), Mainieri et al. (1983), Tomazello Filho et al. (1983), Mainieri & Chimelo (1989), Ilic (1991), and Nardi-Berti & Edlmann Abbate (1992).

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Vessels solitary, in pairs and radial multiples, occasionally up to 7 (Fig. 3), some with gummy or resinous deposits. Intervessel pitting fine (up to c.l 0 /illl), clearly vestured. Fibres mostly very thick-walled. Axial parenchyma scanty paratracheal to vasicentric, aliform and confluent, storied, strands 2-4-celled. Rays uniseriate, rarely biseriate and more rarely up to 3 cells wide (D.jerrea), usually up to c. 15 cells high, storied. Sometimes up to 25 or 35 cells high and axially fused in D. magnifica (MADw 46441).

Pterodon - Fig. 18-21 Perhaps 6 species in Brazil and Bolivia, but critical reappraisal is needed (Polhill 1981 b). Only two species, P. abruptus and P. emarginatus were examined here. Sev­ eral publications contain wood descriptions, including Mainieri & Primo (1971), Detienne & Jacquet (1983), Mainieri et al. (1983), Mainieri & Chimelo (1989), and Nardi-Berti & Edlmann Abbate (1992).

Vessels solitary and in frequent radial multiples, some clusters, some vessels with resinous or gummy deposits. Intervessel pitting fine (up to c. 10 /illl), clearly vestured. Fibres very thick-walled. Axial parenchyma mainly aliform and confluent, strands storied and 2-4-celled. Rays uniseriate, very occasionally biseriate, usually up to 12 cells high, but occasionally up to 22 cells high in P. emarginatus, storied. Prismatic crystals abundant in P. abruptus, not seen in P. emarginatus.

Taralea - Fig. 22-33 About 5 species from tropical South America. Four species were examined here. Taralea oppositijolia is described and illustrated by Anonymous (1981a), as showing storied rays and axial parenchyma as seen in Dipteryx and Pterodon. Detienne et al. (1982) also describe T. oppositijolia (as Dipteryx oppositijolia), not mentioning storeying, although they recognise it in D. odorata and D. punctata. Detienne & J acquet (1983) illustrate D. oppositijolia, which is not storied, presumably from the same wood sample(s). Evidence for storeying in T. oppositijolia is therefore equivocal, although in view of the non-storied nature of all Taralea species examined here (ex­ cept for T. casiquiarensis, see below), it seems most likely that the T. oppositijolia in Anonymous (1981a) is a misidentified Dipteryx or Pterodon.

Vessels solitary, in pairs and radial multiples up to at least 6 in T. oppositijolia. Some vessels with resinous or gummy deposits. Intervessel pitting very fine (up to c. 4/illl). Fibres very thick-walled, most with no lumen. Axial parenchyma scanty paratracheal, aliform to confluent. Most axial parenchyma cells in T. rigida are sclerified (Fig. 27). Axial parenchyma strands 2-4-celled, irregularly or not storied. Rays uniseriate and occasionally biseriate, often less than 20 cells high, but up to 38 cells high in T. oppo­ sitijolia, not storied. Some rays in T. oppositijolia have one row of upright marginal cells. Taralea casiquiarensis differs from the other Taralea species in having much larger intervessel pitting (up to 10.5 /illl) and storied axial parenchyma and rays.

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Fig. 1-4. Dipteryx. All TS. - D. alata, SJRw 50210 (1), D. ferrea, MADw 12465 (2), D. lacunifera (3), D. polyphylla, SJRw 42406 (4). - Scale line is 200 f.UI1 für all.

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Fig. 5 - 8. Dipteryx. All TS. - D. magnifica, MADw 46441 (5), D. micrantha (6), D. panamen­ sis, SJRw 45644 (7), D. punctata, SJRw 34090 (8). - Scale line is 200 ~ for all.

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Fig, 9-13. Dipteryx. -D. rosea, TS (9), D. rosea, TLS (10), D. magnifica, MADw 46441, TLS (11), D. magnifica, SJRw 22606, TLS (12), D. alata, SJRw 50210, TLS (13). - Scale line is 200 /l111 for all.

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Fig. 14-17. Dipteryx. - D. polyphylla, SJRw 42406, TS (14), D. cf.jerrea, MADw 22287, RLS (15), D. alata, SJRw 50210, TLS (16), D. magnifica, MADw 46441, TLS (17). - Scale line is 200 lJIIl for Fig. 14 & 15,50 lJIIl for Fig. 16 & 17.

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Fig. 18-21. Pterodon emarginatus (= P. pubescens). - TS (18), TLS (19), RLS (20), TLS (21). - Scale line is 200 JllIl for Fig. 18-20, 50 JllIl for Fig. 21.

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Fig. 22-25. Taralea casiquiarensis. - TS (22), TLS (23), RLS (24), TLS (25). - Scale line is 200 llID für Fig. 22-24, 50 llID for Fig. 25. - This species is belng reassigned tü Dipteryx by Lewis & Gassün (in press).

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Fig. 26-29. Taralea. T. crassifolia (26, 28, 29), T. rigida, MADw 45218 (27). - TS (26, 27), TLS (28, 29). - Scale line is 200 J..llIl for Fig. 26 & 28, 100 J..llIl for Fig. 27 & 29.

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Fig. 30-33. Taralea oppositijolia, SJRw 23633 (30-32), SJRw 36655 (33). - TS (30), TLS (31), RLS (32), TLS (33). - Scale line is 200 !1Il1 for Fig. 30-32, 100 !1Il1 for Fig. 33.

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COMPARISON OF GENERA AND SPECIES WITHIN DIPTERYGEAE

Dipteryx, Pterodon and Taralea casiquiarensis have a very similar wood anatomy, with short storied rays and storied 2-4-celled axial parenchyma strands. The intervas­ cular pitting is approx. 10 f.llll in horizontal diameter, and vesturing is readily seen on the pit apertures with a light microscope. In TS, the other Taralea species are also similar to Dipteryx and Pterodon, but in TLS differ in having irregularly to non­ storied rays and axial parenchyma, and much smaller intervessel pitting. Pittier (1943) described T. casiquiarensis and explained that he placed the species in Taralea rather than Pterodon on the basis of its larger and fewer leaflets. He did this despite the presence of a winged rachis, which is a feature of Dipteryx (Lewis pers comm.). Pittier's type specimen, which has the same collection number as the wood sampie and presumably came from the same , has been traced to the Field Museum of Natural History in Chicago. This specimen provides ample evidence that the species should have been included in Dipteryx, and this is being rectified by Lewis & Gasson (in press). Size of intervascular pits is a good distinguishing feature in Dipterygeae between Taralea with small pits and Dipteryx and Pterodon with larger ones. I under­ estimated the value of this character in my surveys of Sophoreae and Swartzieae genera (Gasson 1994, 1996).

DIPTERYGEAE AND GENERA IN RELATED TRIBES

Polhill (1981 b) stated that "the tribe (Dipterygeae) certainly comes close to Dal­ bergieae, but the nearest affinity, as suggested by Bentham (1860), seems to be with Monopteryx, an outlier of the Dussia group of Sophoreae." Wood anatomy strongly supports the first statement but not the second.

Dalbergieae - Most genera of Dalbergieae (see Polhill 1981c) have aliform and confluent parenchyma and short regularly storied uniseriate or biseriate rays (e.g., , , , , , , . , Cascaronia, , , , and Grazielodendron). and differ in having wider and taller (i.e. larger) rays, irregu­ larly storied in the former and strongly storied in the latter. Short 1- or 2-seriate storied rays in Dalbergieae are very similar to those in Dipteryx and Pterodon, but not Taralea (except T. casiquiarensis), where they are irregularly storied. Taralea is sometimes referred to Galegeae (Polhill 1981 b) on the basis of its dehiscent fruits, but this tribe is mainly herbaceous and shrubby, and none of the twenty genera listed in Polhill (1981d) are present in RBG Kew's wood or slide collection, so I am currently unable to pursue this.

Sophoreae & Swartzieae - Monopteryx has 10zenge aliform and confluent paren­ chyma, non-storied to irregularly storied rays, 2 cells wide and up to about 35 cells high in M. inpae, and also massive tall and wide rays in addition to smaller ones in M. uaucu (see Gasson 1994, Fig. 91-92). These characters indicate that Monopteryx may not be so closely related to Dipteryx and Pterodon (Dipterygeae) as are most

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Dalbergieae. However, possibly in support of a c10se relationship with Taralea, the intervascular pits are very fine, and although the rays tend to be wider, their arrange­ ment is similar. In Sophoreae very regular storeying of short 1- or 2-seriate rays is found in Sweetia and Luetzelburgia (Gasson 1994, Fig. 23-26), but they have more abundant aliform and confluent parenchyma than most Dipterygeae and Dalbergieae. Some other Sophoreae also have storied rays, which te nd to be slightly wider or taller, e.g., Holocalyx, Uribea, some Ateleia, Pericopsis, and some Dussia. In Swartzieae, Aldina and some Swartzia species are also similar to each other (Gasson 1996) and Dipteryx and Pterodon. Additionally, Aldina (Swartzieae) and Andira (Dalbergieae) are very similar to each other anatornically, and are unresolved in Doyle et al.'s (1997) c1adistic analysis of the rbcL gene. Both have abundant aliform and confluent parenchyma and intervascular pits the size of those in Dipteryx, although the rays are less regularly storied in Andira. In Doyle et al.'s (1997) analysis, Holocalyx is sister to Dipteryx. The wood anat­ omy of these two genera is very similar, although Holocalyx has fine intervascular pitting as found in Taralea. Holocalyx was also discussed in Gasson & Webley (1999), because in Herendeen's (1995) analysis of morphological characters it was found to be basal to four genera of Swartzieae whose wood is very similar (Gasson 1996). All four of these genera (Lecointea, Exostyles, Harleyodendron and Zollernia) have fine intervascular pitting, and the last two have irregularly storied rays. These are features found in Taralea and not Dipteryx. Unfortunately, these observations confuse rather than c1arify the issue. The wood of Taralea and Dipteryx is c1early shown to differ in this paper, but Herendeen's (1995) analysis did not inc1ude any Dipterygeae, and Doyle et al. (1997) did not inc1ude Taralea. There is c1early a limitation to superim­ posing observations of wood anatomy on other authors c1adograms when not all the relevant genera have been inc1uded, and it will be interesting to see where Holocalyx, Dipteryx and Taralea lie in future analyses that also inc1ude Pterodon and all genera of Sophoreae, Swartzieae and Dalbergieae.

Additional wood characters - The discussion above mentions two characters that were not treated in my work on Sophoreae and Swartzieae woods, where 12 anatomi­ cal characters were scored and placed in a c1adistic matrix. The first, intervessel pit size, is highly relevant in this tribe. There is a striking difference in intervessel pit size between Taralea (small pits, c. 4 j.Ull) and Dipteryx and Pterodon (larger pits up to 10.5 j.Ull). Fujii et al. (1994) measured intervessel pits in the Sophora group, where most pits were 6-10 j.Ull, the only ones with smaller pits being Ammodendron and Calpurnia, which had ranges of 4-8 and 4-10 j.Ull, respectively. Examination of the photographs in Gasson (1994, 1997) shows that several genera in Sophoreae and Swartzieae have very fine pitting. The second character, the presence of a few septate fibres in Vatairea, and Hymenolobium (all Dalbergieae), is also of inter­ est. These three genera and Andira differ from the other Dalbergieae especially in ray structure, and are considered by Baretta-Kuipers (1981) to be c10sely allied to Sophoreae. I did not find any septate fibres in Sophoreae or Swartzieae, but they can be easily overlooked when uncommon, or if the fibres are thick-walled.

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CONCLUSIONS

On the basis of wood anatomy, Dipteryx and Pterodon are very similar to each other, and to most genera in Dalbergieae, although there is a wider range ofaxial paren­ chyma patterns in the latter. Taralea differs in two important respects, having irregu­ larly storied rays and fine intervessel pitting, both features that are found in some Sophoreae and Swartzieae, and to a lesser extent Dalbergieae. Monopteryx is quite different from Dipteryx and Pterodon, and not that similar to Taralea.

ACKNOWLEDGEMENTS

I am grateful to Sharon Fisher who cut the seetions. Wood sampies were kindly provided by Regis Miller, Forest Products Laboratory, Madison. All are prefixed MADw or SJRw (see Stern 1988 for details). Gwil Lewis checked the current validity of the scientific names, advised on the taxonomy of the tribe, and obtained the type specimen of Taralea casiquiarensis from Chicago.

REFERENCES

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